Journal of Horticultural Science (1995) 70 (1) 25-34

Temperature affects vegetative growth and flowering of

coffee (Coffea arabica L.).
By J. E. DRINNAN1 and C. M. MENZEL2
1
Queensland Department of Primary Industries, Walkamin Research Station, Walkamin, Qld,
4872, Australia
2
Queensland Department of Primary Industries, Maroochy Horticultural Research Station, P. 0.
Box 5083, Sunshine Coast Mail Centre, Nambour, Qld, 4560, Australia

SUMMARY
The effect of temperature was investigated on the vegetative growth and flowering of
nine coffee cultivars (Catuai Rojo, Catuai, Caturra Amarillo, Caturra Rojo, Catimot,
BMK, SL6, K7 and LB) grown in pots under glass. High day/night temperatures (33°/28°
compared with 18°/13°, 23°/18° and 28°/23°C) accelerated stem extension and node
production. Growth was greatest in cvs K7 and SL6 and lowest in cv. Caturra Rojo.
Prolonged exposure to high temperatures of 33°/28°C accelerated leaf loss and induced a
general decline in tree health. All cultivars showed rapid initial growth during summer
and autumn, and slower growth during winter and spring under short days ( <13 h) and
low irradiances ( <6.8 MJ m-2 d- 1). Growth did not recover in spring and summer at the
end of the experiment. Inflorescences were initiated mainly from April to September
under photoperiods <12 h. More floral buds initiated at 23°/l8°C and 18°/13°C than at
28°/23°C and no floral initiation occurred at 33°/28°C. Inflorescence development took
4-6 weeks at 28°/23° compared with 8-10 weeks at 23°/18° and 12-14 weeks at 18°/13°C.
High temperatures also induced floral malformations. Cvs Catuai Rojo, Caturra Rojo
and Caturra Amarillo had most inflorescences per node and cvs K7, BMK and LB fewest.
Temperatures of 33°/28°C during summer will ensure maximum vegetative growth and
potential number of flowering nodes. Temperatures of 23°/18°C during winter will ensure
healthy and synchronized floral bud development and maximize the number of
inflorescences per node.

COFFEE (Coffee arabica L.) is cultivated in a
range of environments extending 25° north and
south of the equator (Wrigley, 1988) and within
these environments many growth and flowering
patterns are evident. Close to the equator with
no well-defined seasons such as Colombia,
growth flushes and flowering are observed
throughout the year (Morales et al., 1951). In
contrast, in Brazil and Ethiopia with distinct
seasonal climates, there is a single growth and
flowering season (Barros and Maestri, 1972;
Clowes and Allison, 1982). Flowering and
hence fruit ripening extend over several
months even in locations with distinct seasons,
which present major problems for efficient
mechanical harvesting. Compression of the
flowering season would largely overcome these
problems and is the key to mechanical harvest,..
ing. The environmental regimes which favour a
short prolific flowering are not known.
The literature on the influence of environ-
mental factors on growth and flowering of cof.:.
fee under field conditions is confused.
Depending on location, regularity of wet and
dry seasons, temperature, radiation and photo-
period, each may be the dominant factor
controlling the vegetative/reproductive cycle
(Gopal, 1974). The published work also does
not offer an indication of the variation in culti-
var response to changes in climate.
Water supply has a strong effect on flowering
and fruiting in coffee possibly through its
impact on vegetative growth and floral initia-
tion (Morales et al., 1951; Barros and Maestri,
26 Growth and fiowt;ring in coffee
1972; Barros et al., 1978; Alvim, 1985; Cannell,
1985 and Rothfos, 1985). Browning (1973)
reported that shortening the period of vegeta-
tive growth decreased the period of floral initia-
tion. This would be expected to enhance
synchronized flowering. Under the present
growing conditions in North Queensland, floral
initiation occurs from March to September. If
methods could be found to manipulate vegeta-
tive growth so as to compress the period of
floral initiation, synchronizing flowering might
be possible.
Temperature is the other major factor influ-
ence vegetative growth and floral initiation in
coffee (Mes, 1957a; Went, 1957; Nunes et al.,
1968; Gopal and Vasudeva, 1973; Clowes et al.,
1989). Went (1957) found temperatures of 23°/
17°C to be the optimum for vegetative growth.
Nunes et al. (1968) reported depressed growth
above 25°C while Clowes et al. (1989) found
reduced growth above 33°C. High temper-
atures (33°/23° and 30°/17°C) also inhibit floral
development (Mes, 1957a). The optimum tem-
peratures for floral initiation are reported
around 17°C to 23°C, (Mes, 1957a; Gopal and
Vasudeva, 1973). The influence of temperature
on the spread of vegetative growth and floral
initiation is unknown.
This paper reports on the effects of temper-
ature on the vegetative growth and flowering in
nine coffee cultivars. We proposed that low
temperatures would promote uniform floral
initiation, in turn promoting uniform flowering.
Low temperatures would shorten the period of
vegetative growth, decreasing the period of
floral initiation because initiation occurs pro-
gressively along the branch. The low temper-
atures would also promote the onset of floral
initiation (Mes, 1957a; Gopal and Vasudeva,
1973).
MATERIALS AND METHODS
Cultivars and tree culture
The experiment was conducted in Brisbane,
Australia (Lat. 27°S) between January 1989 and
January 1990 using cvs Catuai Rojo, Catuai,
Caturra Amarillo, Caturra Rojo, Catimor,
BMK, SL6, K7 and LB to provide a range in
maturity, yield and vigour. Many of these are
grown commercially in north Queensland.
Seedling trees were grown in 10 1 of peat and
sand (1:1.5) with added fertilizer (Baker, 1957),
in a heated glasshouse with a maximum tem-
perature of 25°C and a minimum night temper-
ature of 15°C. Cultivars SL6, K7, LB and
Catimor were topped to 70 cm, seven months
prior to the experiment and trained to one
sucker during regrowth to match the unpruned
trees.
Treatments
After three years (1 January 1989) plants
were transferred to controlled-temperature
glasshouses maintained at day/night (12 h
days) temperatures of 18°/13°, 23°/18°, 28°/23°
and 33°/28°C. These temperatures regimes
represent the range of temperature likely to be
experienced in coffee growing regions through-
out the world. The plants were grown for 11
months and received 10.4-13.8 h of sunlight
and aµ average total irradiance of 9.2 MJ m-2
d- 1 (45% of outside value). Average daily
vapour pressure deficit ranged from 0.8 to
1.0 kPa.
Measurements
For each of five non-bearing branches per
tree, extension growth, and the numbers of
nodes and inflorescences were recorded at
monthly intervals. As the inflorescences grew,
an average stage of development of the floral
buds on each branch was recorded as well'as the
number of flower buds and any floral abnormal-
ities (see later). The number of leaves absciss-
ing per node were also recorded. Diurnal leaf
water potential [<p1] were recorded over two
separate days using a pressure chamber as
described by Drinnan (1993). Single leaves
were used to determine <p1 of a tree.
Floral growth terminology
The system of classification for floral bud
development devised for this experiment sepa-
rates development into five stages which are
based on unambiguous morphological charac-
ters which are visible to the naked eye
(Figure la).
Stage 1: Buds 1 mm (first visible), green and
covered by bracts. The type of development
cannot be determined at this early stage.
Stage 2: Buds 2-3 mm, dome shaped, covered
by an amber coloured mucilage secreted from
the tip of the inflorescence. The buds can be
determined as potential flowers at this stage.
 J.E. DRINNAN and C. lyL MENZEL 27

(a)

Stage 1 x50 Stage 2 x25

Stage 3 x9 Stage 4 x7 Stage 5 x~

(b)

FIG. 1
a) Five stages of coffee floral development and b) inflorescence with a single star flower x 10. Inflorescences are mounted on
needles. Magnification is shown.
28 Growth and flowering in coffee
Stage 3: Inflorescence 7-9 mm with flower
buds tightly packed together and covered by
mucilage.
Stage 4: Inflorescence ca. 9-12 mm with indi-
vidual flower buds separated and the mucilage
disappearing. Flower buds become lighter
green, cease growth and enter dormancy. The
inflorescences in this stage are sometimes
referred to as fully mature, ready to flower, or in
the critical stage. In this report inflorescences
between stage 4 and less than 5 are still dormant
and only in the inflorescences at stage 5 has
floral bud dormancy been broken.
Stage 5: Open flower or anthesis, results only
after floral bud dormancy is broken.
These stages are approximate as the develop-
ment is continuous rather than in discrete steps
as illustrated here and hence there is qneed to
use half stages to classify bud develop~ent. In
this experiment all the inflorescences develop-
ing on a branch were assigned a stage of
development, and an average stage of develop-
ment calculated for the branch.
The number of star flowers were also
counted (see Figure lb). These flower buds
appear to open prematurely, the petals remain-
ing small and stiff, forming a star shape, the
style and anthers are left exposed. Huxley and
Ismail (1969) reported that a range of inter-
mediate types of flower buds often exist
between a star flower and a normal flower bud.
In some cases the corolla gapes open and the
anthers and style may protrude through the
opening. In this report any of these inter-
mediate stages were recorded as star flowers.
Throughout this paper, the term floral bud
refers to an inflorescence which is developing
(differentiating). Once the floral bud reaches
stage 4, the term flower bud is used because one
or more flower buds will have formed. The term
mature flower bud refers to flower buds which
have fully formed and are in a state of dor-
mancy but which have not reached anthesis.
This period of dormancy before anthesis 1s
referred to as floral bud dormancy.
Analysis
The number of inflorescences, flower buds,
star flowers, dead flower buds, axillary buds and
leaves abscinded were analyzed on a per-node
basis. The increase in the number of nodes,
extension growth and stage of floral develop-
ment were analyzed on"a per branch basis. To
test the effect of cultivar, the data were ana-
lyzed separately for each temperature by one-
way analysis of variance (cultivar only). To test
the effect of temperature over time, the cultivar
means were pooled for each temperature and
the standard error of the gen..~ral mean at each
time presented. Data are the means from six
trees.
RESULTS
Water relations
Leaf water potential decreased with increas-
ing temperature and were 0.2 to 0.5 MPa lower
at midday than at dawn (0600 hours) or sunset
(1800 hours). As the <p1 did not vary significantly
between cultivars only the data for cv. Catuai
Rojo are presented (Table I).
Stem extension and number of nodes
Total shoot growth across cultivars was great-
est at 33°/28°C (13.5 cm per branch) and
declined for each drop in temperature to 5.3 cm
per branch at l8°/l3°C (Table II). Shoot growth
in individual cultivars also decreased ·with
decreasing temperature except in cvs SL6 and
K7 where growth increased slightly at the low-
est temperature (18°/13° compared with 23°/
18°C) (Table II). Growth in these two cultivars
was also good at the highest temperature. Cvs
Catuai and Catuai Rojo grew poorly at low
temperatures but growth improved greatly at
the highest temperature. Cvs LB and Caturra
Rojo were the least respo~sive to increasing
temperature.
Node production generally reflected exten-
sion growth with the greatest increase at 33°/
28°C (6.3 new nodes per branch) and a smaller
increase at 28°/23°C and 23°/l8°C. Node pro-
duction increased slightly at the lowest temper-
ature (Table II). Individual cultivars produced
similar numbers of nodes at each temperature,
the notable exception being cv. Caturra Rojo
which at the highest temperature produced sig-
nificantly fewer nodes than all other cultivars
(Table II). lnternode length was generally
greater during rapid growth and in the vigorous
cultivars.
Most extension growth was produced by the
cultivars pruned before the start of the experi-
ment (cvs SL6, LB, K7 and Catimor). These cul-
tivars averaged 11.1 cm per branch across
 J. E. DRINNAN and C. l\1. MENZEL 29

TABLE I Trees at the highest temperature also devel-

Effect of day/night temperatures on the diurnal leaf water oped soft swollen nodes and prolific axillary
potential (MPa) of cv. Catuai Rojo. Data are the means for six
trees
branching and suckering. Tree growth was
normal and healthy at 23°/l8°C and l8°/13°C.
Time of day
(hours) l8°/13°C 23°/l8°C 28°/23°C 33°/28°C
Leaf drop
0600 0.11 0.22 0.25 0.34 Increased leaf abscission was associated with
1200 0.30 0.47 0.61 0.83 strong vegetative growth at the highest temper-
1800 0.13 0.15 0.20 0.42
atures (33°/28° and 28°/23°C). The average
number of leaves abscinded per node ( ± SE)
temperatures, compared with 5.8 cm extension was 0.66 (0.03) and 0.54 (0.03) at 33°/28° and
growth in the unpruned £¥1tivars. The smallest 28°/23°C compared with 0.18 (0.03) and 0.13
growth rates were produe¢d by the dwarf cvs (0.03) at 23°/18° and l8°/13°C.
Caturra Rojo, Caturra Amarillo and Catuai.
Pruning appeared to be having a strong effect Number of inflorescences
on growth at the start of the experiment as indi- Across cultivars most inflorescences were
cated by the rapid growth of the dwarf cv. Cati- produced at 23°/l8°C followed closely by 18°/
mor. Within the pruned trees however, Catimor l3°C (Table III). Above these temperatures the
grew less than the three vigorous cvs SL6, LB, number of inflorescences produced declined.
K7. Although the pruned trees grew faster, the At 33°/28°C no floral development was
pattern of growth was not affected. observed (Table III). Cv. Caturra Amarillo pro-
duced the highest number of inflorescences per
Pattern of vegetative growth node at all temperatures. Cvs Catimor, LB and
As all cultivars showed similar patterns of K7 produced few inflorescences especially at
growth data were pooled across cultivars. the lowest temperature. In cv. Caturra Rojo the
Growth generally declined from January to number of inflorescences produced increased
May at 23°/18°, 28°/23° and 33°/28° and was uni- at the lowest temperature unlike all other
formly low at l8°/l3°C (Figure 2). Treatment culti vars.
effects were small after June with growth occur-
ring as a series of small flushes. The vegetative Timing of floral initiation
growth cycle of coffee would appear to be For all cultivars, inflorescences were initiated
reasonably continuous throughout the year but from March until September. Initiation was
stronger in summer and autumn. slow at first, reached maximum levels from
April to July and then slowed again. The
Visual observations decline in the number of inflorescences at 28°/
At 28°/23° and especially at 33°/28°C, leaves 23°C after August (Figure 3) was associated
become chlorotic, unusually small, thin and with floral abortion. This pattern of initiation
puckered and the branches thin and elongated. was similar for all cultivars. Cultivars varied
TABLE II
Effect of temperature and cultivar on total extension growth (cm) and number of new nodes per branch (in parenthesis) over
eleven months. Data are the means for five branches over six trees. Temperature means for extension growth (SE = 0.39) and new
nodes (SE= 0.15) are also presented

Cultivar 18°/l3°C 23°/18°C 28°/23°C 33°/28°C

Catimor 6.5 (3.3) 6.9 (3.0) 10.0 (3.7) 15.4 (6.5)

BMK 4.2 (2.0) 4.2 (1.5) 5.8 (2.9) 11.1 (5.3)
Caturra Rojo 2.8 (1.9) 4.3 (2.1) 4.7 (2.6) 8.0 (3.8)
Caturra Amarillo 2.5 (2.1) 2.8 (2.3) 3.6 (2.2) 11.9 (7.0)
LB 8.5 (2.5) 8.7 (2.3) 12.2 (4.3) 12.5 (6.0)
K7 9.8 (3.1) 9.0 (2.5) 12.2 (3.6) 19.0 (7.1)
SL6 8.8 (2.7) 8.0 (2.5) 11.0 (3.3) 18.4 (6.9)
Catuai Rojo 3.0 (2.4) 3.6 (2.4) 7.2 (3.7) 12.1 (7.3)
Catuai 1.3 (1.5) 2.6 (1.4) 6.9 (2.8) 12.8 (6.5)
LSD (P<0.05) 3.3 (1.3) 3.3 (1.3) 3.3 (1.3) 3.3 (1.3)
Mean 5.3 (2.4) 5.6 (2.2) 8.2 (3.2) 13.5 (6.3)

_J
30 Growth and flowering in coffee

:2
-t--' I I I I I I I
c 4
0
E
E
g_ 3
_c
~
0
!......
2
0)
c
0
CJ) 1
c
Q)
-t--'
x
w 0
Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month
2 FIG.
Effect of temperature on the pattern of lateral branch extension during the experi-
ment. Data are the means for all cultivars (36 trees). Vertical bars indicate SE of the
overall mean.

slightly in the timing of initiation: cvs. Catimor, DISCUSSION

Caturra Rojo and Caturra Amarillo initiated Water relations
floral buds earlier than cvs Catuai Rojo and Leaf water potential measurements indicate
Catuai. a more stressful environment with increasing
temperature. However, tree water status at all
Inflorescence development temperature regimes were well within the
Individual inflorescences took approxi- normal diurnal range for fruit trees (Jones et al.,
mately 4-6 weeks at 28°/23°C, 8-10 weeks at
1985).
23°/l8°C and 12-14 weeks at l8°/l3°C to reach
maturity. High rates of inflorescence abscission Stem extension and node number
were observed at 28°/23°C especially during This experiment has shown the large effect of
August and September. Malformed (star) temperature on growth in coffee. The amount
flowers were also observed at 28°/23°C. Cvs of growth (branch extension, new nodes,
Caturra Amarillo, Catuai Rojo and K7 had axillary branches and suckers) increased with
most floral malformations. increasing temperature from 18°/13° to 33°/
TABLE III 28°C. For many cultivars branch extension
Effect of temperature and cultivar on the total number of growth and the increase in the number of nodes
inflorescences initiated per node. Data are the means for five
branches over six trees. Temperature means (SE= 0.14) are was more than three times as great at 33°/28°
also presented. There was no flowering at 33°/28°C compared with l8°/l3°C. Lower optimum tem-
No. of inflorescences per node peratures for vegetative growth have been
established previously by Robinson (1986) 26°/
Cultivar l8°/l3°C 23°/l8°C 28°/23°C 20°c, Willson (1985) 24°/15°C and Went (1957)
Catimor 1.7 2.7 2.0 23°/l 7°C. Willson (1985) reported that the
BMK 2.4 2.4 1.1 photosynthetic rate is reduced above 25°C and
Caturra Rojo 3.5 3.2 1.1
Caturra Amarillo 4.1 5.0 2.0 that leaves are damaged by continual exposure
LB 1.9 2.5 2.0 to temperatures above 30°C. Nunes et al. (1968)
K7 1.7 2.3 1.5 found depressed leaf growth above 25°C.
SL6 1.9 2.9 1.5
Catuai Rojo 3.4 4.9 1.4 In this experiment although total growth was
Catuai 2.8 2.8 0.8 greatest at 33°/28° (mostly due to very high
LSD (P<0.05) 0.7 0.7 0.7
initial rates of growth), continuo~s exposure to
Mean 2.6 3.2 1.5
high temperatures (33°/28° and 28°/23°C to a
 J. E. DRINNAN and C. M. MENZEL 31

Cf)
Q)
u 4
cQ)
u
Cf)

~ 3
0
c
·-D
Q)

a~ 2
ci !......
c 8_
Q)
>
""§
::J
E
0
::J
0
Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month
FIG. 3
Effect of temperature on the pattern of inflorescence initiation during the experiment.
Floral initiation was inhibited at 33/28°C. Data are the means for all cultivars (36 trees).
Vertical bars indicate SE of the overall mean.

lesser extent) led to reduced rates of growth
and deteriorating plant health. Leaves became
chlorotic, small and short lived. Nodes became
swollen and malformed and branches became
long, thin and spindly. Therefore the optimum
temperature for tree health ( <28°/23°C) is
lower than for branch extension growth. and
similar to optimum temperatures reported by
Went (1957), Willson (1985) and Robinson
(1986).
Growth rates of individual cultivars indicates
that cvs K7 and SL6 are tolerant of low and high
temperatures, cvs Catuai and Catuai Rojo are
also tolerant of high temperatures but cold sen-
sitive while extension growth in cvs LB and
Caturra Rojo is fairly unresponsive to changing
temperatures. Because of the strong associ-
ation between the amount of vegetative growth
and yield in coffee, these findings may influence
the choice of cultivar for particular locations.
For example cv. C~tuai or Catuai RojcYmay be
selected for low-elevation sites with high mean
temperatures while cvs K7 and SL6 may be
selected for higher elevated sites with cooler
mean temperatures. The low number of nodes
produced by cv. Caturra Rojo at the highest
temperature would limit the yield potential of
this cultivar at this temperature.

I
D
14 15 ""•
E
D 13 ·~hot<iper;od /•-• • J
0
2...__...
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Q)
Q_
0
+--'
12
11
.-------. • ....__
lrradiance~ / .-·/
10

0 5
_c -----.~.-
Q_ 10
0
Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month
FIG. 4
Average monthly photoperiod and irradiance inside the glasshouse during the experiment.
32 Growth and fi0,wering in coffee
The pattern of vegetative growth found in
this study is similar to patterns found by Tau-
send (unpublished) in Hawaii, Barros and
Maestri (1972) in Brazil and Clowes and Alli-
son (1982) in Ethiopia. The pattern appears to
be related to changes in photoperiod or
irradiance rather than temperature. The
general decline in growth from January to May
corresponded with a decline in photoperiod
and irradiance (Figure 4). A reduction in veg-
etative growth has been observed under short
days by Franco (1940), Piringer and Borthwick
(1955) and Went (1957) and in low irradiances
by Rayner (1946). A decline in growth at this
time has also been related to competition with
fruit growth by Mcfarlane (1949), Beaumont
and Fukunaga (1958) and Cannell (1971), but
the decline in this experiment was not associ-
ated with cropping. The reason for the poor
growth in spring and early summer under a
steadily increasing photoperiod and irradiance
is not known.
Number of inflorescences
This experiment has shown the large effect of
temperature on floral development in coffee.
Floral initiation is promoted below 28°/23°C
and eliminated at 33°/28°C. Development of
inflorescences is faster at 28°/23°C compared
with 23°/l8°C and 18°/13°C but this leads to
floral malformations and inflorescence ~bscis­
sion. Mes (1957a) found that most floral buds
were produced at 23°/17°C with none at 30°/
23°C or 30°/17°C. In other studies Kumar (1979)
found trees at 27°/17° developed the most floral
buds. Gopal and Vasudeva (1973) found floral
initiation was favoured by temperatures
around 22°-23°C.
Floral initiation in individual cultivars indi-
cates that cv. Caturra Amarillo is suitable for a
range of temperatures while cvs Catimor, LB
and K7 appear less suited to low temperatures
and that cv. Caturra Rojo is most suited to low
temperatures. These findings will influence the
choice of cultivar for maximizing potential
yield at particular locations.
The similar pattern of floral initiation
between temperatures suggests that floral
initiation is controlled by an environmental fac-
tor other than temperature, possibly photo-
period. Inflorescences appeared only when the
photoperiod was < 12-13 h and reached a
maximum rate under the short days of mid-
winter and then declined again as day length
increased (Figure 4). Franco (1940) and Pir-
inger and Borthwick (1955) found a critical
photoperiod for floral initiation in coffee of
13 h.
The timing of floral initiation in each of the
cultivars corresponds well with the field matu-
rity times found by Winston and O'Farrell
(1987 and 1988). Cvs Catuai and Catuai Rojo
are late maturing and cvs Catimor, Caturra
Rojo and Caturra Amarillo mature early.
For synchronous flowering, a short period of
floral initiation would be advantageous. At 28°/
23°C, floral development is rapid, so that within
3.5 months most of the inflorescences have
developed compared with five to six months at
18°/13°C (Figure 3). However, at this temper-
ature many star flowers are formed. A temper-
ature regime of 23°/l8°C appears to be the best
compromise between healthy and synchron-
ized floral bud development. More flower buds
per inflorescence are also formed at 23°/18°C
with 1.2 ( ± 0.09) compareq with 0.15 ( ± 0.09) at
28°/23°C pooled across cultivars. Floral malfor-
mations appear to be related to temperature
and not water deficits as indicated by changes in
cp 1 across temperature. Differences between cul-
tivars suggest some genetic control as well.
High temperatures and water stress have been
suggested as responsible for the formation of
star flowers in earlier studies (Mes, 1957b;
Went, 1957; Alvim, 1958; Huxley and Ismail,
1969; Kumar, 1982).
CONCLUSION
Temperature strongly affected vegetative
and reproductive growth in coffee. High tem-
peratures (33°/28° and 28°/23° compared with
23°/18° or l8°/l3°C) increased the amount of
growth but did not influence the timing of
flushes. Flushing was generally stronger at the
start of the experiment during summer under a
photoperiod >13 hand irradiance >9 MJ m- 2
d- 1, declined during winter, but did not recover
in the following spring and summer. Cultivars
did not vary significantly in the timing of veg-
etative flushes. As yield is related to the amount
of extension growth in the previous season
(Beaumont and Fukunaga, 1958; Montoya et
al., 1961; Cannell, 1971), temperatures of about
33°/28°C during summer should produce the
great~st yields.
 J.E. DRINNAN and C. M. MENZEL 33

Floral initiation did not occur above 28°C or
when· the photbperiod was longer than 13 h.
Floral initiation generally occurred as vegeta-
tive growth was declining. Temperatures around
z3°/l8°C during winter will ensure healthy and
synchronous floral bud development and maxi-
mize the number of inflorescences per node.
Cultivars did not vary significantly in the pattern
of floral initiation, which would have a strong
influence on the synchronization of flowering.
The cultivars with the best potential for high
yields will be those which produce high numbers
of nodes during summer and high numbers of
inflorescences per node during winter, for
example cv. Catuai Rojo.
The influence of photoperiod on the vegeta-
tive and reproductive growth cycles is not
resolved. In south Brazil (Lat. 25°S) floral
initiation occurred only during May and June
(Alvim, 1958) presumably due to the large
seasonal variation in photoperiod. This would
greatly assist synchronized flowering.
The assistance of Dr W. Slater with manu-
script preparation and C. Howitt in analysis and
interpretation of the data was most appre-
ciated. The authors are also most grateful to
John Mansfield for helping in data collection
and to the University of Queensland glasshouse
staff for help in looking after the trees. Finan-
cial support from the Rural Industries
Research and Development Corporation
(RIRDC) is gratefully acknowledged.

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(Accepted 14 June 1994)